Pteridospermatophyta
The term Pteridospermatophyta refers to several distinct polyphyletic groups of extinct seed-bearing plants. The earliest fossil evidence for plants of this type is the genus Elkinsia of the late Devonian age. They flourished particularly during the Carboniferous and Permian periods. Pteridosperms declined during the Mesozoic Era and had mostly disappeared by the end of the Cretaceous Period, though some pteridosperm-like plants seem to have survived into Eocene times, based on fossil finds in Tasmania.
The concept of pteridosperms goes back to the late 19th century when palaeobotanists came to realise that many Carboniferous fossils resembling fern fronds had anatomical features more reminiscent of the modern-day seed plants, the cycads. In 1899 the German palaeobotanist Henry Potonié coined the term "Cycadofilices" for such fossils, suggesting that they were a group of non-seed plants intermediate between the ferns and cycads. Shortly afterwards, the British palaeobotanists Frank Oliver and Dukinfield Henry Scott made the critical discovery that some of these fronds were associated with seeds that had identical and very distinctive glandular hairs, and concluded that both fronds and seeds belonged to the same plant. Soon, additional evidence came to light suggesting that seeds were also attached to the Carboniferous fern-like fronds Dicksonites, Neuropteris and Aneimites. Initially it was still thought that they were "transitional fossils" intermediate between the ferns and cycads, and especially in the English-speaking world they were referred to as "seed ferns" or "pteridosperms". Today, despite being regarded by most palaeobotanists as only distantly related to ferns, these spurious names have nonetheless established themselves. Nowadays, four orders of Palaeozoic seed plants tend to be referred to as pteridosperms: Lyginopteridales, Medullosales, Callistophytales and Peltaspermales.
Their discovery attracted considerable attention at the time, as the pteridosperms were the first extinct group of vascular plants to be identified solely from the fossil record. In the 19th century the Carboniferous Period was often referred to as the "Age of Ferns" but these discoveries during the first decade of the 20th century made it clear that the "Age of Pteridosperms" was perhaps a better description.
During the 20th century the concept of pteridosperms was expanded to include various Mesozoic groups of seed plants with fern-like fronds, such as the Corystospermaceae. Some palaeobotanists also included seed plant groups with entire leaves such as the Glossopteridales and Gigantopteridales, which was stretching the concept. In the context of modern phylogenetic models, the groups often referred to as pteridosperms appear to be liberally spread across a range of clades, and many palaeobotanists today would regard pteridosperms as little more than a paraphyletic 'grade-group' with no common lineage. One of the few characters that may unify the group is that the ovules were borne in a :wiktionary:cupule|cupule, a group of enclosing branches, but this has not been confirmed for all "pteridosperm" groups.
With regard to the enduring value of the division, many palaeobotanists still use the pteridosperm grouping in an informal sense to refer to the seed plants that are not angiosperms, coniferoids, ginkgophytes or cycadophytes. This is particularly useful for extinct seed plant groups whose systematic relationships remain speculative, as they can be classified as pteridosperms with no valid implications being made as to their systematic affinities. Also, from a purely curatorial perspective the term pteridosperms is a useful shorthand for describing the fern-like fronds that were probably produced by seed plants, which are commonly found in many Palaeozoic and Mesozoic fossil floras.
Phylogeny
An alternative phylogeny of spermatophytes based on the work by Novíkov & Barabaš-Krasni 2015 with plant taxon authors from Anderson, Anderson & Cleal 2007 showing the relationship of extinct clades.Taxonomy
- Family ?†Avatiaceae Anderson & Anderson 2003
- Order ?†Alexiales Anderson & Anderson 2003
- * Family †Alexiaceae Anderson & Anderson 2003
- Order ?†Hamshawviales Anderson & Anderson 2003
- * Family †Hamshawviaceae Anderson & Anderson 2003
- Order ?†Hexapterospermales Doweld 2001
- * Family †Colpospermaceae Doweld 2001
- * Family †Hexapterospermaceae Doweld 2001 nom cons.
- Order ?†Hlatimbiales Anderson & Anderson 2003
- * Family †Hlatimbiaceae Anderson & Anderson 2003
- Order ?†Matatiellales Anderson & Anderson 2003
- * Family †Matatiellaceae Anderson & Anderson 2003
- Order ?†Petriellales Taylor et al. 1994
- * Family †Petriellaceae Taylor, del Fueyo & Taylor 1994
- * Family †Kannaskoppiaceae Anderson & Anderson 2003
- Order †Umkomasiales Doweld 2001
- * Family †Angaropeltidaceae Doweld 2001
- * Family †Umkomasiaceae Petriella 1981 Stockey & Rothwel nom. illeg.; Zuberiaceae Němejc 1968]
- Class ?†Arberiopsida Doweld 2001
- * Order †Dicranophyllales Meyen 1984 emend. Anderson, Anderson & Cleal 2007
- ** Family †Dicranophyllaceae Archangelsky & Cúneo 1990
- * Order †Aberiales Meyen 1984
- ** Family †Schmeissneriaceae Zhou 1997
- ** Family †Arberiaceae Anderson & Anderson 1985
- Class ?†Axelrodiopsida Anderson & Anderson
- * Order †Axelrodiales Anderson & Anderson 2007
- ** Family †Zamiostrobacea Anderson & Anderson 2007
- ** Family †Axelrodiaceae Anderson & Anderson 2007
- Class †Moresnetiopsida Doweld 2001
- * Order †Calamopityales Němejc 1963
- ** Family †Calamopityaceae Scott 1909
- * Order †Tetrastichiales Němejc 1968
- ** Family †Tetrastichiaceae Němejc 1968
- * Order †Pullarithecales Doweld 1998
- ** Family †Gnetopsidaceae Doweld 2001
- ** Family †Pullarithecaceae Doweld 1998
- ** Family †Calathiopsidaceae Doweld 2001
- ** Family †Austrocalyxaceae Vega & Archangelsky 2001
- * Order †Moresnetiales Doweld 2001
- ** Family †Eurystomataceae Long 1975
- ** Family †Eospermatacesidae Long 1975
- ** Family †Moresnetiaceae Němejc 1963 emend. Anderson, Anderson & Cleal 2007
- Class †Lyginopteridopsida Novák 1961 emend. Anderson, Anderson & Cleal 2007
- * Order †Lyginopteridales Havlena 1961
- ** Family †Lyginopteridaceae Potonie 1900 emend. Anderson, Anderson & Cleal 2007
- Class †Pachytestopsida Doweld 2001
- * Order †Codonospermales Doweld 2001
- ** Family †Codonospermaceae Doweld 2001 emend. Anderson, Anderson & Cleal 2007
- * Order †Pachytestales Doweld 2001 Sterzel 1896; Whittleseyaceae Remy & Remy 1959; Callipteridaceae Corsin ex Wagner 1965; Callipteridiaceae; Callipteraceae; Protoblechnidaceae]
- ** Family †Alethopteridaceae Corsin 1960 emend. Anderson, Anderson & Cleal 2007
- ** Family †Cyclopteridaceae Corsin ex Wagner 1964
- Class †Callistophytopsida
- * Order †Callistophytales Rothwell 1981 emend. Anderson, Anderson & Cleal 2007
- ** Family †Cornucarpaceae Doweld 2001
- ** Family †Callistophytaceae Stidd & Hall 1970 nom. cons.
- Class †Peltaspermopsida Doweld 2001
- * Order †Sporophyllitales Doweld 2001
- ** Family †Sporophyllitaceae Doweld 2001
- ** Family †Leuthardtiaceae Doweld 2001
- * Order †Trichopityales Doweld 2001
- ** Family †Trichopityaceae Němejc 1968
- * Order †Peltaspermales Taylor 1981
- ** Family †Cardiolepidaceae Meyen 1977
- ** Family †Autuniaceae Doweld 2001
- ** Family †Peltaspermaceae Pilger & Melchoir 1954
- Class †Phasmatocycadopsida Doweld 2001
- * Order †Phasmatocycadales Doweld 2001
- ** Family †Phasmatocycadaceae Doweld 2001
- * Order †Gigantopteridales Li & Yao 1983
- ** Family †Emplectopteridaceae Wagner 1967
- ** Family †Gigantopteridaceae Koidzumi 1936
- Class †Pentoxylopsida Pant ex Doweld 2001
- * Order †Pentoxylales Pilger & Melchior 1954
- ** Family †Lindthecaceae Anderson & Anderson 2003
- ** Family †Pentoxylaceae Pilger & Melchior 1954
- Class †Dictyopteridiopsida Doweld 2001
- * Order †Dictyopteridiales McLoughlin ex Doweld 2001
- ** Family †Breyteniaceae Doweld 2001
- ** Family †Dictyopteridiaceae Rigby 1978
- * Order †Lidgettoniales Doweld 2001
- ** Family †Denkaniaceae Doweld 2001
- ** Family †Parthaceae Doweld 2001
- ** Family †Lidgettoniaceae Anderson & Anderson 1985
- * Order †Rigbyales Doweld 2001 Zimmermann 1930 nom. rej.]
- Class †Cycadeoideopsida Scott 1923
- * Order †Fredlindiales Anderson & Anderson 2003
- ** Family †Fredlindiaceae Anderson & Anderson 2003
- * Order †Cycadeoideales Berry 1920
- ** Family †Varderkloeftiaceae Anderson & Anderson
- ** Family †Laurozamitiaceae Anderson & Anderson
- ** Family †Benneticarpaceae Anderson & Anderson
- ** Family †Westersheimiaceae Němejc 1968
- ** Family †Sturianthaceae Doweld 2001
- ** Family †Williamsoniaceae Nathorst 1943
- ** Family †Williamsoniellaceae Nakai 1943
- ** Family †Cycadeoideaceae R. Br. ex Wieland 1908
- Class †Caytoniopsida Thomas ex Frenguelli 1946
- * Order †Caytoniales Gothan 1932
- ** Family †Caytoniaceae Kräusel 1926